Processing of color- and noncolor-coded signals in the gourami retina. III. Ganglion cells.

The dynamics of intracellular responses from ganglion cells, as well as that of spike discharges, were studied with the stimulus regimens and analytic procedures identical to those used to study the dynamics of the responses from horizontal and amacrine cells (,). The stimuli used were large fields of red and green light given as a pulsatile input or modulation about a mean luminance by a white-noise signal. Spike discharges evoked by a white-noise stimulus were analyzed in exactly the same manner as that used for analysis of analog responses. The canonical nature of kernels allowed us to correlate the first- and second-order components in a spike train with those of the intracellular responses from horizontal, amacrine, and ganglion cells. Both red and green stimuli given alone in darkness produced noncolor-coded responses from all ganglion cells. In the case of some cells, steady red illumination changed the polarity or waveform of the response to green light. Color-coded ganglions responded only to simultaneous color contrast. Nonlinearities recovered from intracellular responses, and spike discharges were similar to those found in responses from amacrine cells and were of two types, one characteristic of the C amacrine cells and the other characteristic of the N amacrine cells. The first-order kernels of most ganglion cells could be divided into two basic types, biphasic and triphasic. The combination of kernels of these two basic types with different polarities can produce a wide range of responses. Addition of two types of second-order nonlinearity could render color coding in this relatively simple retina as an extremely complex process. Color information appeared to be represented by the polarity, as well as the waveform, of the first-order kernel. The response dynamics is a means of transmission of color-coded information. Second-order components carry information about changes around a mean luminance regardless of the color of an input. Some spike discharges produced a well-defined cross-kernel between red and green inputs to show that a particular time sequence of red and green stimuli was detected by the retinal neuron network. The similarity between signatures of second-order kernels for both amacrine and ganglion cells indicates that signals undergo a minimal transformation in the temporal domain when they are transmitted from amacrine to ganglion cells and then transformed into a spike train. Under our experimental conditions, a single spike train carried simultaneously information about red and green inputs, as well as about linear and nonlinear components. In addition, the spike train also carries a cross-talk component. A spike train is a carrier of multiple signals. Conversely, many types of information in a stimulus are independently encoded into a spike train.